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New skeletal elements of the recently described endemic giant
anseriform Garganornis ballmanni Meijer, 2014 are presented,
coming from the type-area of the Gargano and from Scontrone,
southern and central Italy, respectively. The new remains
represent the first bird remains found at Scontrone so far, and
another shared element between these two loc...
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Citations
... Moreover, fossils suggest that extinct spurred birds tended to be flightless and giantsized (e.g. Andrews, 1901;Leguat, 1708;Livezey, 1996;Pavia 2017;Williams, 2015;Worthy, 2001)-a pattern congruent with our results. ...
Birds are a remarkable example of how sexual selection can produce diverse ornaments and behaviours. Specialised fighting structures like deer's antlers, in contrast, are mostly absent among birds. Here, we investigated if the birds’ costly mode of locomotion—powered flight—helps explain the scarcity of weapons among members of this clade. Our simulations of flight energetics predicted that the cost of bony spurs—a specialised avian weapon—should increase with time spent flying. Bayesian phylogenetic comparative analyses using a global spur dataset corroborated this prediction. First, extant species with flight-efficient wings (which presumably fly more frequently) tend to have fewer or no bony spurs. Second, this association likely arose because flying more leads to more frequent evolutionary loss of spurs. Together, these findings suggest that, much like pneumatic bones, absence of weaponry may be another feature of the avian body plan that allows birds to efficiently explore the aerial habitat.
... These two localities have been grouped in the same palaeobioprovince, the Apulo-Abruzzi area . The majority of the different fissures of Gargano pertain to a timespan between the late Miocene and the Pliocene and have yielded a considerably rich fossil fauna over the past decades (Freudenthal, 1971(Freudenthal, , 1972(Freudenthal, , 1976(Freudenthal, , 1985Ballmann, 1976;Leinders, 1984;Abbazzi et al., 1996;Delfino et al., 2007;Masini et al., 2010Masini and Fanfani, 2013;Freudenthal et al., 2013;Villier and Carnevale, 2013;Meijer, 2014;Van Der Geer, 2014;Savorelli and Masini, 2016;Pavia et al., 2017;Pavia, 2013); younger, Early Pleistocene sites, known as Pirro Nord are also known in Gargano (for the reptiles see, among others, Delfino and Bailon, 2000;Blain et al., 2019). Scontrone is older (Tortonian) and has yielded a poorer, though still unique faunal assemblage, comprising also endemic mammals and birds with peculiar morphologies (Rustioni et al., 1992;Mazza and Rustioni, 1996, 2011Savorelli et al., 2017;Meijer, 2014;Van Der Geer, 2014;Pavia et al., 2017). ...
... The majority of the different fissures of Gargano pertain to a timespan between the late Miocene and the Pliocene and have yielded a considerably rich fossil fauna over the past decades (Freudenthal, 1971(Freudenthal, , 1972(Freudenthal, , 1976(Freudenthal, , 1985Ballmann, 1976;Leinders, 1984;Abbazzi et al., 1996;Delfino et al., 2007;Masini et al., 2010Masini and Fanfani, 2013;Freudenthal et al., 2013;Villier and Carnevale, 2013;Meijer, 2014;Van Der Geer, 2014;Savorelli and Masini, 2016;Pavia et al., 2017;Pavia, 2013); younger, Early Pleistocene sites, known as Pirro Nord are also known in Gargano (for the reptiles see, among others, Delfino and Bailon, 2000;Blain et al., 2019). Scontrone is older (Tortonian) and has yielded a poorer, though still unique faunal assemblage, comprising also endemic mammals and birds with peculiar morphologies (Rustioni et al., 1992;Mazza and Rustioni, 1996, 2011Savorelli et al., 2017;Meijer, 2014;Van Der Geer, 2014;Pavia et al., 2017). ...
... Although not so well known as the nearby Gargano palaeoisland, Scontrone is characterized by an astonishing endemic fauna, comprising, among others, large erinaceids, the peculiar hoplitomerycids, and large non-flying birds (Van Der Geer, 2014;Pavia et al., 2017;Savorelli et al., 2017). Several of these insular taxa have congeneric relatives only in Gargano and nowhere else in the world, such as the iconic large erinaceid Deinogalerix Freudenthal, 1972(Savorelli et al., 2017. ...
We here describe a small turtle assemblage originating from the early Tortonian (late Miocene) palaeoisland of Scontrone, central Italy, a locality previously known mostly for its endemic mammals and giant birds, which were otherwise shared only with the Gargano localities, another fossiferous area belonging to the same palaeobioprovince. The fossil turtle remains from Scontrone are referred to the geoemydid Mauremys sp. and a so far unidentified large-sized testudinid. The biogeographic origins of the Scontrone insular chelonians are discussed. The Scontrone geoemydid adds to the known occurrences of Mauremys in the late Miocene of the Mediterranean. The Scontrone large tortoise represents the oldest known Mediterranean insular testudinid, predating significantly the well-known Quaternary endemic island tortoises of the area.
... The term 'giant bird' is used to refer to all avialans (extant and extinct) that have evolved into large-sized forms, standing out from other members of their respective clades. The most common use of this term in the literature refers to terrestrial and flightless birds (e.g., Baskin, 1995;Buffetaut and Le Loeuff, 1998;Worthy et al., 2016;Angst and Buffetaut, 2017;Pavia et al., 2017) that move in a cursorial or graviportal way and are large in size, according to a human scale. Ecologically, these kinds of giant birds are diverse in habitats and diets (Fig. 1A). ...
Throughout the evolutionary history of Avialae, several members of this clade have evolved into giant forms, in different time periods and ecological contexts. In Europe, the first birds that show this condition, the Gargantuaviidae, occur during the Late Cretaceous (late Campanian–early Maastrichtian), but it is during the Paleogene when more groups evolve large forms. However, until now, there was no record of any giant bird during the late Maastrichtian of Europe, close to the K/Pg boundary. Here we describe a cervical vertebra (MPZ 2019/264) from Beranuy (Huesca, NE Spain), which is the first
fossil evidence of a giant bird from the late Maastrichtian of Europe, within Chron C29r. The vertebra displays some features, such as a well-marked heterocoelous articulation, lateral pneumatic foramina, ventral carotid processes, and a low neural spine, that support its inclusion within the clade Ornithuromorpha. This phylogenetic assignment is supported by two cladistic analyses. The vertebra is clearly different from the one assigned to Gargantuavis, meaning that it belonged to a distinct taxon. Although the kinship between these two taxa of giant birds is still unclear, this finding demonstrates that
large-sized birds were part of the ecological communities of the Ibero-Armorican island from the late Campanian to the Late Maastrichtian, being present during the last hundreds of thousands of years prior to the K/Pg extinction event.
... Modena INTRODUCTION Insular environments are well known to represent laboratories of evolution, characterised by rapid species diversification, high levels of endemism, minimum or maximum size extremes, and peculiar morphologies (among others, Van der Geer et al., 2010). Documented cases of unique taxa, as well as extreme forms characterised by both dwarfism and gigantism are known in the fossil record already since the Mesozoic, with principal examples the Cretaceous dwarf dinosaurs of the Haţeg Island in modern day Romania (Benton et al., 2010), the Neogene and Quaternary flightless birds of numerous oceanic islands (Steadman, 2006;Worthy et al., 2016;Pavia et al., 2017), the Neogene and Quaternary giant rodents of the Mediterranean islands ( Van der Geer et al., 2010), the Neogene and Quaternary dwarf hippos, deers, and peculiar "artiodactyls" of the Mediterranean islands (Palombo et al., 2006(Palombo et al., , 2012b(Palombo et al., , 2013; Van der Geer et al., 2010), the small carnivorans from the Mediterranean islands Abbazzi et al., 2005;Van der Geer et al., 2010), the Sardinian Plio-Pleistocene macaque (Rook & O'Higgins, 2005;Zoboli et al., 2016), and, most principally, the Quaternary dwarf elephants and mammoths of the Mediterranean islands, the Philippines, and Indonesia (Major, 1883;Van der Geer et al., 2010;Herridge & Lister, 2012;Palombo et al., 2012a;Zoboli et al., 2018). Insular extinct Cenozoic reptile taxa are not so well known as their co-occurring mammal species; however, there are known interesting cases of large-sized lacertids and giant tortoises in the late Neogene and Quaternary of the Mediterranean (Leith Adams, 1877; Böhme & Zammit-Maempel, 1982;Bailon et al., 2014;Rhodin et al., 2015;Luján et al., 2017), as well as peculiar terrestrial crocodylians, large iguanians, and bizarre chelonians in the remote islands of Oceania (Pregill & Dye, 1989;Mead et al., 2002;Pregill & Worthy, 2003;Pregill & Steadman, 2004;White et al., 2010). ...
We herein describe Sardophis elaphoides Georgalis & Delfino n. gen. n. sp., a new snake taxon from the early Pleistocene of Monte Tuttavista VI, Sardinia, Italy. Sardophis elaphoides Georgalis & Delfino n. gen. n. sp. possesses a distinct vertebral anatomy and
is diagnosed by a unique combination of features. The new Sardinian taxon is further compared in detail and differentiated from all extant European and North African snake species. Although the affinities of Sardophis elaphoides Georgalis & Delfino n. gen. n. sp. with colubroids are clear, its more inclusive relationships within that clade cannot be resolved with certainty. Being an insular form, Sardophis elaphoides Georgalis & Delfino n. gen. n. sp. adds significantly to our so far poor knowledge of island endemic snakes. An overview of the fossil record of snakes from the Mediterranean islands is provided. The new species increases the number of reptile taxa that went extinct in Sardinia during the late Neogene and Quaternary.
... Similarly, osteological analyses indicate substantial phenotypic differentiation between prehistoric (NZ) versus modern (Australian) populations (figure 2). Specifically, the extinct NZ lineage exhibited classic morphological signs of the 'island rule' effect [43,[46][47][48][49] (figure 2b,c), suggesting a relatively terrestrial life history compared to that of the extant Australian lineage. The distinctive large body size, elongated legs, proportionally short robust wings relative to legs, and substantially increased body mass of the extinct prehistoric swans suggest that this lineage was on an evolutionary pathway towards flightlessness [44,48]. ...
... Our study provides a rare example of a putatively flightreduced and perhaps semi-terrestrial swan lineage. Indeed, only two suggested flightless swan taxa (both extinct), one of uncertain affinity within Anatidae, have previously been recorded in the fossil record [44,57,58] compared to numerous flight-reduced or flightless ducks and geese [44,49,59]. While substantial changes in relative body size are commonplace in insular island species [10,49,[59][60][61], they have not previously been clearly characterized within Cygnus (e.g. ...
... Indeed, only two suggested flightless swan taxa (both extinct), one of uncertain affinity within Anatidae, have previously been recorded in the fossil record [44,57,58] compared to numerous flight-reduced or flightless ducks and geese [44,49,59]. While substantial changes in relative body size are commonplace in insular island species [10,49,[59][60][61], they have not previously been clearly characterized within Cygnus (e.g. [44,57]). ...
Prehistoric human impacts on megafaunal populations have dramatically reshaped ecosystems worldwide. However, the effects of human exploitation on smaller species, such as anatids (ducks, geese, and swans) are less clear. In this study we apply ancient DNA and osteological approaches to reassess the history of Australasia’s iconic black swans (Cygnus atratus) including the palaeo-behaviour of prehistoric populations. Our study shows that at the time of human colonization, New Zealand housed a genetically, morpho-logically, and potentially ecologically distinct swan lineage (C. sumnerensis, Poūwa), divergent from modern (Australian) C. atratus. Morphological analyses indicate C. sumnerensis exhibited classic signs of the ‘island rule’ effect, being larger, and likely flight-reduced compared to C. atratus. Our research reveals sudden extinction and replacement events within this anatid species complex, coinciding with recent human colonization of New Zealand. This research highlights the role of anthropogenic processes in rapidly reshaping island ecosystems and raises new questions for avian conservation, ecosystem re-wilding, and de-extinction. © 2017 The Author(s) Published by the Royal Society. All rights reserved.
... The loss of flight ability, which has occurred numerous times in the evolutionary history of birds (Raikow 1985, Feduccia 1999, Livezey 2003, may lead to morphological specialization, recent studies on which have provided novel insights into avian biology. Examples include use of the wing as a weapon (Longrich and Olson 2011, Hume and Steel 2013, Williams 2015a, Pavia et al. 2017, neuroanatomical modifications (Iwaniuk et al. 2004, 2009, Smith and Clarke 2012, and structural and histological modifications of bones (Habib and Ruff 2008, Habib 2010, Smith and Clarke 2014, De Mendoza and Tambussi 2015. ...
Flight ability has been lost many times in the family Anatidae (ducks, geese, swans, and allies), and this provides unique insights into the morphological and ecological evolution of the family. Although ~15 fossil anatids have been reported to be flightless or possibly so, there has not been an established criterion that is widely applicable to assessing flight ability in fossil anatids. In this study, discriminant rules for the presence–absence of flight ability were constructed by linear discriminant analysis (LDA) based on 7 skeletal measurements in 93 modern anatids in order to set a basis for the inference of flight ability in fossil anatids. Model selection for LDA was conducted by a high-dimensional modification of Akaike’s Information Criterion, and selected models discriminated the volant and flightless groups with only one misclassification (Tachyeres patachonicus). Flight abilities of fossil anatids were assessed by the constructed rules, supplemented by resampling experiments that were designed to assess the uncertainty in estimating skeletal proportions of fossil anatids in the absence of associated skeletons. The flightless condition was strongly supported for Cnemiornis spp., Branta rhuax, Hawaiian moa-nalos, Chenonetta finschi, Anas chathamica, Chendytes spp., Shiriyanetta hasegawai, Cayaoa bruneti, and the "Annaka Short-winged Swan," whereas the volant condition was supported for Mergus milleneri and Bambolinetta lignitifila. Results were ambiguous for Branta hylobadistes and Anas marecula. The constructed rules can easily be applied to new observations in the future, although limitations in the inference of ecological traits in fossil species from morphological measurements, including the risk of extrapolations, should be appreciated.
... In recent years, newly discovered Italian localities provided information that partially bridges the chasm of knowledge. Until recently, the vertebrate fossil record from the late Miocene of Italy have consisted solely of endemic faunal complexes mainly from two isolated areas: the Apulo-Abruzzi paleobioprovince (e.g., Freudenthal, 1971;Mazza and Rustioni, 2011;Delfino and Rossi, 2013;Masini et al., 2013;Meijer, 2013;Patacca et al., 2013;Pavia, 2013;Villier and Carnevale, 2013;Savorelli et al., 2016;Pavia et al., 2017) and the Tusco-Sardinian paleobioprovince (e.g., Engesser, 1989;Abbazzi et al., 2008a;Rook et al., 2011). However, some of the newly discovered Italian fossil localities yield nonendemic faunal assemblages. ...
The systematic analysis of more than 20,000 fossils (Vertebrata and Mollusca), recovered from the post-evaporitic Messinian (5.41-5.33 Ma) succession of Moncucco Torinese (NW Italy), resulted in the identification of 90 vertebrate and 65 mollusk taxa that provide additional information about the paleoecological context and the paleoenvironmental settings of NW Italy slightly before the Mio-Pliocene boundary. Our analyses indicate a landscape dominated by open woodlands within a mosaic environment also including closed canopy forests, grasslands, rocky outcrops and limited water edges. The wide spectrum of habitats may have had a prominent role in determining the high paleobiodiversity observed in the paleocommunity of Moncucco Torinese. Slight variations in the abundances of the most common rodent species over the investigated succession are probably related to local changes in the paleolandscape. From a paleoclimatic point of view, the overall information provided by the fauna indicates mesic conditions in a subtropical climate, which is also consistent with the interpretation derived from paleobotanical and sedimentological analyses for the latest Messinian of Northern Italy.
This chapter gives a concise overview of the faunas of Gargano. Then, a comprehensive treatment is provided of the history of discoveries, biozones, or faunal units and the peculiarities and evolutionary aspects of individual endemic species or lineages. The palaeo‐island Gargano is now part of mainland Italy. During the Late Miocene and Early Pliocene, it was an island, harbouring a highly endemic fauna, including five‐horned deer, giant gymnures, giant pikas, and several rodent lineages. The mid‐Pliocene flooding of the area led to the extinction of this peculiar fauna. After the area emerged again and became connected to Italy in the Early Pleistocene, it was colonised by a balanced mainland fauna. From the Early Pleistocene on, the region was interested by a general uplifting, resulting in the regression and continentalisation of the area.
